Characters useful in distinguishing larvae of Popillia japonica and other introduced scarabaeidae from native species

Historic, archived document

Do not assume content reflects current
scientific knowledge, policies, or practices.

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CIRCULAR No. 334 DECEMBER 1934

UNITED STATES DEPARTMENT OF AGRICULTURE
WASHINGTON, D C.

CHARACTERS USEFUL IN DISTINGUISHING LARVAE
OF POPILLIA JAPONICA AND OTHER INTRODUCED
SCARABAEIDAE FROM NATIVE SPECIES

By Rosert J. Sim, chief illustrator, Division of Japanese and Asiatic Beetles,
Bureau of Entomology and Plant Quarantine

CONTENTS

Page Page
UT trom Ch OM ese eee ne ees 2 a et 1 | Phyllophaga ephilida (Say)-----------.------- 7
Popilliayaponica Newme. 225 lo eee Suey Llophagatristisy (ia bs) se ee ee 8
Anomala orientalis Waterh___.____.__________ 4 | Polyphylla variolosa (Hentz)___-_--__--_---_- 8
AMO ALA DIN OLOLa Gye ee ee AY Ochrosidia t2llissa. (Burm) ee ee 9
Strigoderma arboricola (Fab.)__-_______- a mele 4 | Aphonus castaneus (Melsh.)_______.-__-____- 9
Pachystethus lucicola (Fab.)-----.-.---_--___- Di eA LOT CLES SALUTES) CEIAD») eae aoe ee eae 9
PEAT OLUE DUN CECE (as) eee ee eee ee Det | es COLUN SERIE A Clas) pee ees re ey 10
Macrodactylus subspinosus (Fab.)_-_____-__- a Bri eA DRONE ANAGs Claeys ter te lee ees 10
Autoserica castanea Arrow__----_---_-_------ 6 | Osmoderma eremicola Knoch_____------------ 10
SERED SUNILTS MLC WIS se os ee ne ee Culp Al phogiusOssors (ls) seen eee ee ee 11
SCRICALDATAUCLT, OSVP Eee ee se eae ogee 6 TF OTISUDCTOSIES CHADS) = ay os ee ee 11
PIU DLOLALES SP) wee os nes eet aes P/O (ai eblecusiakend© happiness sie ees ee 12
DCH ELOTY TIS aoe a tes es TS eg eI 7

INTRODUCTION

This circular is intended to present in the simplest way the
characters most convenient to use in distinguishing the larvae of
Popilia japonica and other introduced species from native species
which are frequently associated with them. Twenty-four species
met with in the eastern part of the United States are briefly described
and figured. Special stress is placed upon the epipharyngeal struc-
tures and the characters of the tenth or last abdominal segment
(posterior end) of the grub.t Where expedient, brief notes on the
behavior and habitat are given. .

For preliminary field determinations involving only general ap-
pearance and anal characters, a 10-power hand lens is used. For
more careful examination in the laboratory, a low-power binocular
microscope is required, together with a watch glass, forceps, and a
needle ground to a cutting edge.

The familiar term “ white grub” is usually applied to the larvae
of May beetles (Phyllophaga spp.), but so far as general appearance
goes it could be used for any of the scarabaeid species. These may be
described as short, more or less stout, blunt-ended grubs of a creamy

1In this circular the nomenclature, with the exceptions of raster, sensillae, sclerotized,
and sensory eminence, is that given in the following publication: HaAyrs, W.
MORPHOLOGY, TAXONOMY, AND BIOLOGY OF LARVAL SCARABAEOIDEA. [1]. Biol. Monographs
Veco aoe, 1929.

82234°—34——_1 1

De CIRCULAR 334, U. S. DEPARTMENT OF AGRICULTURE

or whitish color with darker (yellowish or brownish) heads and
slight vestiture. Three pairs of short legs are attached to the
anterior fourth of the larva on its ventral side. In its usual posi-
tion—when not crawling—the grub lies curled up in the form of a
horseshoe. The surface is more or less shining, but is deeply wrin-
kled transversely. There are short spines or setae dorsally, and on
the end and ventral surface of the terminal posterior segment is a
definite arrangement of spines and hairs. This group of spines and
setae, as seen from beneath, is called the raster, and offers the most
obvious characters for field determinations. At or near the center
of this group there are, In many species, a number of dark, stout
spines arranged variously as a transverse row, 2 longitudinal rows,
or 2 curved or diverging rows. These conspicuously modified
spines are, in some species, sufficient for immediate identification
without reference to other characters.

The next most useful features are on the head of the grub—the
color and “surface finish” of the head capsule and the structures
of the epipharynx.

In most species the characters of the head capsule, epipharynx,
and raster are sufficient for determination, but in a few the claws
may be examined to advantage. In any case, the length and general
proportions of a grub should be noted; some species are more slender
than others. The normal habitat may be a decaying tree, soil about
shrub and tree roots, leaf humus in woodland, the turf of lawns,
fields, and pastures, high or low sandy soil, the moist humus of
open swamps, ete. Any such distinct condition of environment is
generally helpful in making determinations.

The epipharynx is the inner (or under) surface of the labrum,
although some of its structures extend back on the inner wall of
the clypeus. In order most successfully to examine the epipharyn-
geal structures, an incision should be made with a very small knife
along the clypeal suture which extends transversely between the
bases of the mandibles. The entire mass comprising the labrum
and clypeus is then turned over and placed in a watch glass of
alcohol. If fine particles of foreign matter obscure the structures
when the whole is examined under a binocular microscope, they may
be removed by careful manipulation of a small pointed brush. For
comparison with the drawings, the specimen should be examined
with the apex of the labrum pointing away from the observer (the
base of the clypeus toward him). Looked at thus, it will be seen
that somewhat below the apex there is a raised area or eminence
terminating in a sclerotized point or several large fixed spines. This
small mound could be called the sensory eminence, for on it are
various arrangements of sensillae. These appear as small dark
circles, usually in two sizes. At the base of the labrum are two
blackish sclerotized plates called the tormae. These extend inward
from the sides and may end in acute points or more rarely may be
fused with each other, thus forming a narrow dark band across the
base of the labrum. The shape of each torma should be noted as a
good character in determination. Between the apices of the tormae
are several special structures more or less specific in character. At

CHARACTERS DISTINGUISHING LARVAE OF POPILLIA, ETC. 3

the apex of the right torma (at left in the drawings) are two rather
large sclerotized objects, usually brown in color. These are the
sclerotized plate and the sense cone. There may be a few small
sensillae and groups of fine sensory hairs near these. Near the apex
of the left torma are usually several small sensillae and lkewise
groups of sensory hairs. In many species there is an elongate group
of larger forked sensory hairs above the apex of the left torma.

With the exception of a rather large, concave, smooth median
space, the surface of the epipharynx is sparsely armed with articu-
lated spines and hairs. These are largest toward the central space
and gradually decrease in size toward the lateral margins of the
epipharynx. Just inside each lateral margin there is usually a series
of marginal striae.

The entire margin is more or less armed with coarse articulated
spines of varying lengths—always greater in the apical region.

As may be seen in the drawings, all the structures are more or
less useful in making determinations. Special attention should be
paid to the head capsule. Its surface may be glossy or opaque;
glabrous, hairy, or setose; smooth, punctured, or reticulated. In
color it may be some tone of dull yellow, brown, or blackish. The
characters of the head capsule are described in each species. The
structures mentioned are shown in the explanatory drawings (fig. 1).
The descriptions are by no means complete, as it has seemed unde-
sirable to confuse the observer with discussions of the antennae,
mandibles, maxillae, labium, legs, etc. Aside from those of the
epipharynx, only the more easily discernible characters are noted.
The length measurements given are approximate, having been made
as well as possible from alcoholic specimens. The term ‘“ form
medium ” is applied to those species which are neither unusually
stout as in Osmoderma, nor somewhat slender as in Macrodactylus.

POPILLIA JAPONICA NEWM.

(Introduced)

Length 23 mm. Form medium.

Width of head 3 mm.

Surface of head: Smooth, shining. Epicranial stem a fine, dark, impressed
line. Epicranial arms not conspicuous. Front with a short, vague, longi-
tudinal, median impression in apical third. At each side of this a row
of five punctures diverging toward middle bend of epicranial arm.

Color of head: Pale dull yellow.

Epipharynx (fig. 1) : Sensory eminence with a strong, angled sclerotized plate
in right side. Several sensillae of two sizes near base of the group of
three large, fixed teeth. Right torma long, slender, slightly sinuate; left
torma very slender, with apical third curved toward apex of labrum, and
with a rather large basal expansion.

Raster (fig. 6) : Numerous coarse, rather long, scattered, brown, hooked spines.
Medially two conspicuous, divergent rows of shorter, straight spines in
V form; 6 or 7 spines in each row. At sides and end of tenth segment
numerous rather long, yellowish hairs.

Anal slit: Transverse, arcuate.

Vestiture: Entire grub with rather long scattered brown hairs. Dorsal con-
vexities of first six abdominal segments clothed with fine short brown spines.

Habitat: Soil under turf.

Remarks: The distinct V-like arrangement of short dark spines of the raster
are sufficient to identify this species.

4 CIRCULAR 334, U. S. DEPARTMENT OF AGRICULTURE

ANOMALA ORIENTALIS WATERH.
(Introduced)

Length 25 mm. Form medium.

Width of head 3 mm.

Surface of head: Shining, shallowly reticulated and with several brown setae.
Epicranial stem in anterior half a fine impressed line with raised edges.
Epicranial arms barely apparent. Front with two paramedian setose
punctures near clypeal suture. Two transversely placed setose punctures
below middle curve of epicranial arm.

Color of head: Brownish yellow vaguely clouded with brown.

Epipharynx (fig. 2): A vague sclerotized plate on the right below apex of
labrum. Sensory eminence with a distinct, wide, arcuate, transverse,
sclerotized plate. In the curve of this, several sensillae and 3 or 4 large
fixed teeth.

Raster (fig. 6): Subanal convexity with scattered, coarse, rather long, hooked
spines. Below this two paramedian rows of very short stout spines, about
14 in each; these rows parallel or slightly divergent at anal end. Outside
of these, on each side, two sparse irregular rows of hooked spines smaller
than those of subanal convexity. Sides and apex of terminal segment with
numerous, rather long, brown hairs.

Anal slit: Transverse, slightly arcuate.

Vestiture: Generally as in Popillia japonica, but short dorsal spines slightly
more conspicuous.

Habitat: Soil of certain nurseries, lawns, and golf courses.

Remarks: In Popillia japonica, Anomala orientalis, A. binotata, Pachystethus
lucicola, and Strigoderma arboricola the epipharyngeal structures are simi-
lar; the slight differences in the sensory eminence, tormae, and sensillae
shown in the drawings should be carefully noted.

ANOMALA BINOTATA GYLL.

Length 23 mm. Form as in Popillia japonica.

Width of head 3.5 mm.

Surface of head: As in A. orientalis, but distinctly and finely reticulate and
frontal punctures less apparent.

Color of head: Pale dull yellow vaguely clouded with darker.

Epipharynx (fig. 2): Apparently without sclerotized plate near apex of
labrum. The transverse arcuate plate partly encircling the sensillae and
four fixed teeth is very narrow.

Raster (fig. 6): Similar to that of A. orientalis, but the two paramedian rows
of short spines always clearly divergent at anal end.

Anal slit: Transverse, arcuate.

Vestiture: Generally as in Popillia japonica, but short dorsal spines slightly
finer and closer.

Habitat: Sandy fields and lawns.

Remarks: The reticulate head capsule, the finer frontal punctures, and the
definitely divergent lines of spines of the raster characterize this species.

STRIGODERMA ARBORICOLA (Fab.)

Length 23 mm. Form as in Popillia japonica.

Width of head 3 mm.

Surface of head: Shining, feebly and finely granulate. Epicranial stem very
feebly impressed. Frontal punctures 3 or 4 between clypeal suture and
curve of epicranial arm. Front without median impression.

Color of head: Pale dull yellow.

Epipharynx (fig. 2): Generally similar to that of Anomala orientalis, but
sclerotized plate on right of sensory eminence distinct and smaller. The
darkened area including the sensillae of sensory eminence less broad.
Apex of left torma longer.

Raster (fig. 6): Similar to that of Popillia japonica, but the two paramedian
rows of short spines more irregular, longer, and nearly parallel.

Anal slit: Transverse, arcuate. :

Vestiture: In general as in Popillia japonica.

CHARACTERS DISTINGUISHING LARVAE OF POPILLIA, ETC. 5

Habitat: Usually waste, sandy fields and dunes.

Remarks: Difficult to distinguish from some specimens of Popillia japonica,
but the punctation of the head and the characters of the raster are usually
sufficient.

PACHYSTETHUS LUCICOLA (Fab.)

Length 23 mm. Form of Popillia japonica.

Width of head 2.5 mm.

Surface of head: Shining, finely reticulate. Occipital half of epicranial stem
an impressed line with feebly raised edges. Front uneven; two paramedian
setose punctures near clypeal suture; above these, on each side, 2 to 4
transversely placed towards middle curve of epicranial arm.

Color of head: Pale dull yellow.

Hpivharynx (fig. 2): Very similar to that of Strigoderma arboricola, but sen-
sory hairs of epigusta finer and more numerous. No slender sclerotized
plate at right of epigusta.

Raster (fig. 6): Similar of that of Strigoderma arboricola, but the two para-
median rows of pointed spines more regular, diverging throughout, with
the individual spines longer, eight spines in each row.

Anal slit: Transverse, arcuate.

Vestiture: As in Popillia japonica, but small dorsal spines finer.

Habitat: Under turf of pastures, lawns, and fields.

Remarks: This grub is usually yellowish white throughout instead of being
darkened by the earthy contents in the ninth and tenth segments. The
V of the raster is longer than in P. japonica and the two sides are less
divergent.

PELIDNOTA PUNCTATA (L.)

Length 40 mm. Form rather stout.

Width of head 6 mm,

Surface of head: Mostly glabrous, faintly glossy, with extremely fine, confused,
shallow wrinkles. Epicranial stem narrow, dark, impressed. Epicranial
arms very fine, raised. Front rather finely, irregularly punctured ante-
riorly, the larger punctures with fine yellow setae.

Color of head: A rather bright brownish ocher, darker anteriorly.

Epipharynx (fig. 3); Semicircular in outline. Sensory eminence with a curved
sclerotized plate, in center of which is a large short fixed spine. Scat-
tered below plate are several sensillae. Between inner ends of tormae
is a short angular chitinous bar; below this, several sensillae. Spines
along lateral margins of epipharnyx very short.

Raster (fig. 7): Hairs and spines all rather fine and sparse, of a bright rufous
color.

Anal slit: Transverse; sinuate as seen from end of grub.

Vestiture: Dorsal surface of abdominal segments (exclusive of the tenth)
with very fine, short, and rather sparse brown hairs interspersed with
a few long hairs.

Habitat: The decaying but still firm portions of old stumps, frequently maple
and poplar. Usually considerable numbers of these grubs, in various
sizes, are found in a single stump.

Remarks: Feeding, as it does, upon decaying wood which is not much dis-
colored, this grub is not usually as dark posteriorly as are most scara-
baeid larvae.

MACRODACTYLUS SUBSPINOSUS (Fab.)

Length 15 mm. Form elongate, rather slender.

Width of head 2 mm. ;
Surface of head: Smooth, shining. Epicranial stem a fine depressed line.
Epicranial arms not apparent. Front with six scattered punctures.

Color of head: Dull yellow clouded with rufous. e

Epipharynx (fig. 2): With a small apical lobe. Sensory eminence with an
obleng sclerotized plate towards the right side. Below this a triangular
plate at each end of the row of five sensillae. Four short, blunt, fixed
teeth at proximal margin of eminence. The right torma angulately bent
at apical third.

6 CIRCULAR 334, U. S. DEPARTMENT OF AGRICULTURE

Raster (fig. 6): Two paramedian rows of short, stout, well-separated spines
D to 7 on each side. Outside of these are widely spaced, heavy, hooked
spines about twice as long. On the sides of the terminal segment are
a few long scattered hairs. The end is clothed densely with somewhat
shorter hairs.

Anal slit: Y-shaped.

Vestiture: Entire grub sparsely clothed with long yellow hairs. Dorsal con- —

vexities of first five abdominal segments with numerous short, brown
spines.
Habitat: Soil of nurseries, lawns, and fields. ;
Remarks: The rather slender form and the anal characters distinguish this
common grub,

AUTOSERICA CASTANEA ARROW
(Introduced)

Length 15 mm. Form rather slender.

Width of head 2.5 mm.

Surface of head: Shining, mostly glabrous, but with two short, fine setae on
each side of vertex, and one in a circular depression just below epicranial
branch on each side of front. Epicranial arches very faintly discernible.

Color of head: Pale ocher (dull yellow).

Epipharynx (fig. 4): Curved sclerotized bar below continuous and joining
sclerotized plate on right of sensory eminence. Long spines at each side
of sehsory eminence more numerous and more closely placed. Left torma
slender but rather short.

Raster (fig. 6): The transverse curved row of brown spines composed of 18
to 20. The short spines anterior to this (below in drawing) small and
numerous.

Anal slit: Longitudinal, forking dorsad.

Vestiture: The first six abdominal segments evenly covered dorsally with
very short brownish setae interspersed with a few long fine hairs.

Habitat: Soil of fields, gardens, and lawns of infested areas.

Remarks: This rather slender grub is very active and restless. The very
short claw of the leg is characteristic.

SERICA SIMILIS LEWIS
(Introduced)

Length 15 mm. Form rather slender.

Width of head 2 mm.

Surface of head: As in Auwtoserica castaned.

Color of head: As in A. castanea.

Epipharynx (fig. 4): Curved sclerotized bar below apex not apparent in the
specimens examined. Long spines at each side of sensory eminence large
and more numerous than in S. parallela. Left torma rather long—longer
and more slender than in A. castanea.

Raster (fig. 6): Transverse, curved row of brown spines composed of 14 to
18. The short spines anterior to this few in number and very coarse—
much fewer and coarser than in A. castanea.

Anal slit: As in A. castanea.

Vestiture: Similar to that of A. castanea.

Habitat: Soil of limited areas on Long Island.

Remarks: Only rather undeveloped third-instar larvae have been available
for examination. See remarks under Serica paraliela.

SERICA PARALLELA CSY.

Length 15 mm. Form somewhat slender.
Width of head 2.3 mm.
Surface of head: As in Autoserica castanea.
Color of head: As in A. castanea.
Epipharynx (fig. 4) : Curved sclerotized bar below apex interrupted at middle.
Long spines at each side of sensory eminence few and distant. Left
torma short and stout.

CHARACTERS DISTINGUISHING LARVAE OF POPILLIA, ETC. 7

Raster (fig. 6): Characters intermediate between those of A. castanea and
Serica similis.

Anal slit: AS in A. castanea.

Vestiture: Similar to that of A. castanea, but the short dorsal setae darker,
finer, and slightly less dense.

Habitat: Sandy soil, usually in oak woods and thickets of coastal plains.

Remarks: The three sericine larvae herein described are similar in general
characteristics. The slight differences in the epipharynges, rastelrs, and
claws of third legs should be noticed carefully.

DIPLOTAXIS SP.

Length 22 mm. Form somewhat slender.

Width of head 2.5 mm.

Surface of head: Shining, shallowly reticulated. Groove of epicranial stem
with narrowly raised edges. Epicranial arms deeply impressed, nearly
straight at middle. Front with three large setose punctures on each side—
one near middle, one just below middle of epicranial arm, and one near
antennal end of epicranial arm.

Color of head: Rather light dull yellow.

Epipharynx (fig. 2): Labrum of the usual suborbicular shape with a vaguely
indicated median lobe. Sensory eminence with 6 or 7 larger and 3 smaller
sensillae; a rather vague sclerotized plate on right side, and 3 fixed teeth.
Right torma long and rather slender; left torma shorter, heavy at base
and tapering toward apex. Sense cone and chitinous plate of epigusta
small.

Raster (fig. 6): On each side a conspicuous blackish patch of pointed spines,
longer and more dense toward anal slit. Lateral and apical hairs of
terminal segment rather dense but short.

Anal slit: Y-shaped.

Vestiture: About as in Popillia japonica.

Habitat: So far as observed, found in sandy soil.

Remarks: The species shown is from Long Island and is probably D. atlantis
Fall, which is common there.

DICHELONYX SP.

Length 18 mm. Form slender, as in Macrodactylus subspinosus.

Width of head 2 mm.

Surface of head: Shining, slightly or not reticulated. Anterior half of epicranial
stem a darkened groove with raised edges. Epicranial arms very slightly
impressed. Frent with 4 setose punctures—2 paramedian close to clypeal
suture and 1 on each side near middle of epicranial arm.

Color of head: Pale dull yellow.

Hpipharynx (fig. 2): Outline of labrum somewhat angular; a strong middle
lobe with a large convexity on each side; the lateral margins subserrate
and with short, coarse spines. Sensory eminence with several sensillae
and 2 or 8 large fixed teeth; it is flanked by a vague, oblong sclerotized
plate near each side. Tormae nearly meeting above epigusta, their tips
obscured by sensory hairs and a large sclerotized plate.

Raster (fig. 6) : A median and terminal group of coarse rather long spines, well
spaced; at each side an elongate, longitudinal band of very small, short,
out-pointing spines. End of terminal segment with a tuft of long hairs
at each side.

Anal slit: Y-shaped.

Vestiture: As usual, but inconspicuous.

Habitat: This grub has been found in the soil in nurseries.

Remarks: Easily recognized by the elongate lateral patches of short, small
spines of the raster. The species figured is from northern New Jersey
and is probably D. elongata (Fab.).

PHYLLOPHAGA EPHILIDA (Say)

Length 28 mm. Form medium.

Width of head 3.5 mm.

Surface of head: Opaque, smooth. Epicranial stem a fine, impressed, dark
line. Arms not conspicuous. Two very small punctures near each arm on
front.

8 CIRCULAR 334, U. S. DEPARTMENT OF AGRICULTURE

Color of head: Dull yellow.

Epipharynx (fig. 4) : Apex obtusely pointed, with margins sinuous. Marginal
striae long and distinct. An oblique row of five vague striae near each
side of sensory eminence. The eminence has 2 pale sclerotized plates, an
arcuate row of 6 large sensillae, and an irregular group of dark fixed
teeth. Between and below the tormae are many sensillae and sensory hairs.

Raster (fig. 6): Two long nearly parallel rows of small, short spines. Out-
side of these, numerous much longer, brown, hooked spines merging into
still longer, fine yellow hairs at sides and apex. The small paramedian
spines are not closely placed, and the two rows converge slightly toward
the anal end.

Anal slit: Broadly Y-shaped.

Vestiture: A few scattered long pale hairs throughout. Dorsal convexities
of first Six abdominal segments with fine, pale, inconspicuous, evenly spaced
spines.

Habitat: Soil under turf in pastures, lawns, ete.

Remarks: Of the numerous Phyllophaga species, this and tristis are the only
ones that have been reared at Moorestown, N. J.

PHYLLOPHAGA TRISTIS (Fab.)

Length 25 mm. Form medium.

Width of head 3.5 mm.

Surface of head: Opaque, smooth. Epicranial stem a fine dark line impressed
in anterior half. Arms not conspicuous. Front with a distinct subcircular
impression at each side of middle near clypeal suture and two small punec-
tures below middle of epicranial arm.

Color of head: Light brownish yellow.

Epipharynx (fig. 4): Similar to that of P. ephilida, but without submarginal
striae, with only one sclerotized plate (right) in sensory eminence, and
fewer sensillae on epigusta.

Raster (fig. 6) : Two well-separated, arcuate rows of short brown spines. Out-
side of these a few longer, scattered, hooked spines. Sides of terminal
segment with sparse, rather long brown hairs, those of apex somewhat
shorter.

Anal slit: A broad Y.

Vestiture: Practically as in ephilida.

Habitat: Soil under turf of lawns, pastures, and open grassy spots in wood-
land.

Remarks: The two paramedian rows of spines of the raster resemble paren-
theses. These are distinctive. While the epipharynges of most Phyllo-
phaga grubs have submarginal striae, no trace of these could be detected
in the several tristis specimens available.

POLYPHYLLA VARIOLOSA (Hentz)

Length 40 mm. Form medium,

Width of head 6 mm.

Surface of head: Opaque because of fine reticulation. Anterior part of front
more distinctly roughened. Epicranial stem impressed, the anterior third
with raised margins. Epicranial arms searcely apparent. Three seta-
bearing punctures on each side of front. Clypeus and labrum rough.

Color of head: Yellowish brown with two vague, longitudinal, lighter bands.

Epipharynx (fig. 5): To the right of the group of fixed spines on the sensory
eminence are a dozen or more small articulated spines. The right side of
the median bare area is bordered with numerous fine hairs. There are no
submarginal striae.

Raster (fig. 7): Numerous short hooked spines, with a short, distinct, double
row of short straight spines medially. Each side densely clothed with
longer brown hairs.

Anal slit: Transverse, clearly sinuate.

Vestiture: First five abdominal segments dorsally clothed rather densely and
evenly with short brown setae. Sides glabrous, venter with numerous
rather long brown hairs. '

CHARACTERS DISTINGUISHING LARVAE OF POPILLIA, ETC. 9

Habitat: Sandy soil of the east coast States. These larvae feed upon plant
roots, are sometimes found in the turf of golf courses, and have been known
to kill privet hedge and small maple trees.

Remarks: Distinguished from similar sized Phyllophaga grubs by the short
but dense brown vestiture of the back and the short double row of spines
in the center of the raster.

OCHROSIDIA VILLOSA (Burm.)

Length 24 mm. Form medium or slightly stout.

Width of head 4 mm.

Surface of head: Shining, shallowly but sharply reticulate. Upper part of
epicranial stem a fine impressed line with broadly swollen margins. Front
with a large roundish impression at each side of middle, each with a setose
puncture; a transverse row of three small punctures above and outside
these impressions. Rather numerous small punctures near clypeal suture.

Color of head: Brownish yellow clouded with darker.

Epipharynx (fig. 4): Sensory eminence with a single large, dark, sclerotized
plate with a rounded apex, in the left side of which is a deep notch. The
sense cone on the epigusta is rather small.

Raster (fig. 7): A sparse group of coarse, long, hooked brown spines becoming
larger towards anal slit. Apex of terminal segment somewhat truncate
and clothed with yellow hairs of moderate length.

Anal slit: Transverse, arcuate.

Vestiture: Normal, i.e., a general very sparse distribution of long slender hairs,
and rather densely placed, very short brown dorsal spines.

Habitat: Common and often harmful in the turf of lawns and golf courses.

Remarks: Slightly larger and stouter than the Popillia grub and may be dis-
tinguished from it in the field by lack of V in the raster.

APHONUS CASTANEUS (Melsh.)

Length 28 mm. [orm rather stout.

Width of head 3.5 mm.

Surface of head: Shining, but (except at extreme sides) hairy and roughened,
with rather close and coarse setose punctures. Epicranial stem a distinct
impressed line. Epicranial arms obscured by the general punctation.

Color of head: Chestnut brown, slightly lighter at sides.

Epipharynx (fig. 4): Sensory eminence with a single large, dark, sclerotized
plate. The apex comes to a broadly obtuse point directed towards the
large median: spineless area. On the right margin a single large articulate
tooth. Sense cone on epigusta large and elongate.

Raster (fig. 7) : An extensive patch of brown, hooked spines, similar to that of
Ochrosidia villosa but with the spines more numerous and noticeably
smaller. End of abdomen rounded—not slightly truncate as in Ochrosidia.

Anal slit: Distinctly arcuate.

Vestiture: Fairly numerous long hairs throughout; the small dorsal spines
more hairlike and longer than usual.

Habitat: Sandy soil. Rather frequent in turf of golf courses of the coastal
plains.

Remarks: The dark, rough, hairy head immediately distinguishes this grub
from any others herein described. ‘

XYLORYCTES SATYRUS (Fab.)

Length 55 mm. Form stout.

Width of head 7 mm. ‘

Surface of head: Shining, glabrous, coarsely punctured throughout, vaguely
impressed on each side of front. Epicranial stem impressed at top of
vertex.

Color of head: Brownish black.

Epipharynx (fig. 5): The striking characters are the large blunt beaklike
sclerotized projection of the sensory eminence and the T-shaped apex of
the left torma.

Raster (fig. 7): As shown, without special structures.

82234°—34

2

10 CIRCULAR 334, U. S. DEPARTMENT OF AGRICULTURE

Anal slit: Transverse, slightly sinuate at ends.

Vestiture: Surface mostly glabrous; first five abdominal segments dorsally
with very short, sparse, dark-brown spines, interspersed with very few
long brown hairs.

Habitat: The humus under dead leaf carpet of deciduous woods. Usually 2
or 3 grubs may be found within 3 square yards.

Remarks: The prothoracic sclerotized plate is rufous brown and conspicuous.
The large distinct spiracles are darker brown. This is the largest of the
northern scarabaeid grubs.

COTINIS NITIDA (L.)

Length 38 mm. Form rather stout, blunt, flattened beneath.

Width of head 5 mm.

Surface of head: Mostly glabrous, feebly shining, finely roughened with irreg-
ular, impressed reticulation. Epicranial stem and arms showing distinctly
as impressed light lines, the former seeming to continue downward as a
median furrow in upper half of front. Surface of front irregular; a large
puncture or pit below the middle of each epicranial arm.

Color of head: Chestnut brown, blackish near Clypeal suture.

Epipharynx (fig. 3) : Three-lobed; the marginal depression each side of median
lobe with a small darkish mark. Sensory eminence not pronounced. About
12 short, fixed spines in a strongly curved transverse row. No distinct
sclerotized plate in median lobe. Right torma extended basally, below.

Raster (fig. 7) : Numerous fine hairs of medium length, becoming longer at the
sides and end of the terminal segment. Medially a rather long double row
of very short, stout spines quite irregularly placed.

Anal slit: Short, broadly arcuate.

Vestiture: Practically the entire grub clothed rather evenly, but inconspicuously,
with fine brown hairs, those on the dorsal surface much shorter than the
hairs of the venter.

Habitat: In soil under manure or decaying vegetation in open places.

Remarks: This grub, like most of the Cetoniinae, is a “ back-crawler.” The
prothoracic sclerotized plates are rather large, well defined, yellowish brown.
The legs are short and brownish yellow.

EUPHORIA INDA (L.)

Length 35 mm. Form short, stout, blunt.

Width of head 4 mm.

Surface of head: Mostly glabrous, shining, very feebly reticulate. Furrow of
epicranial stem extending downward on front between arms. A large
puncture just below middle bend of epicranial arm on each side of front.

Color of head: Brownish yellow.

Epipharynx (fig. 3) : Similar to that of Cotinis, but with a long, narrow, curved,
rather feeble sclerotized plate above the row of fixed spines of the sensory
eminence. A definite but thin sclerotized plate above sense cone. Few, if
any, spines below apex of left torma.

Raster (fig. 7) : Similar to that of Cotinis, but vestiture more sparse and without
any special median spines.

Anal slit: Transverse, nearly straight.

Vestiture: Very slight, limited to a narrow band of fine, short, stiff hairs (1 or
2 hairs wide) on each convex fold; those of the dorsal surface very short,
the very sparse lateral and ventral hairs much longer.

Habitat: Usually in and under old horse manure.

Remarks: A “ back-crawler.” One favored habitat is the old mixed sawdust

and horse manure about an abandoned sawmill in the woods.

OSMODERMA EREMICOLA KNOCH

Length 45 mm. Form stout, distinctly larger posteriorly and blunt at the ends.

Width of head 6.5 mm.

Surface of head: Nearly glabrous; rather shallowly, irregularly, but distinctly
reticulated. Epicranial stem dark and deeply impressed. Epicranial arms
showing distinctly as narrow light-yellow lines. Front irregularly rough-
ened. Mandibles short, stout, and very irregular in surface.

CHARACTERS DISTINGUISHING LARVAE OF POPILLIA, ETC. ua

Color of head: Vertex yellowish brown; front darker brown.

Epipharynx (fig. 3): Three lobed, the depression in the outline at each side
of median lobe with a small dark mark. The sensory eminence not pro-
nounced but broad and with a slightly curved transverse row of about 12
short fixed spines. Above this are several small sensillae. An elongated
sclerotized plate in the right side of the median lobe.

Raster (fig. 7): A large area of sparse, very short blackish spines medially,
rather numerous but scattered longish dark hairs laterally and apically.

Anal slit: Transverse, arcuate.

Vestiture: Entire surface very sparsely clothed with fine but stiff dark-brown
hairs, short to medium in length. This vestiture noticeable to the touch
but not conspicuous to the naked eye.

Habitat: The much-decayed interior of dying tree trunks. Frequently many
of varying sizes are found in one tree.

Remarks: The prothoracie sclerotized plates are large, sharply defined, and
yellowish brown: the spiracles, dark brown.

APHODIUS FOSSOR (L.)

Length 10 mm. Form short, stout, dilated posteriorly.

Width of head 3 mm.

Surface of head: Opaque, very finely closely reticulated. Epicranial stem
impressed, margins raised in anterior third. Front with longitudinal
furrow in upper fourth; a vague vertical depression on each side just
above clypeal suture.

Color of head: Rather dark bright brown.

Epipharynx (fig. 3): Definitely three-lobed. Sensory eminence with a long
transverse line of minute sensillae, below which is a semicircle of short
toothlike spines continuing laterally to base of labrum. Central bare
area with a tuft of fine hairs above and to right of middle, which has a
dark line extending from base. A band of fine hairs between apices
of the short tormae.

Raster (fig. 6): Numerous very short, stout blackish spines rather evenly
placed. On each side a few slender hairs.

Anal slit: Transverse, arcuate; three glabrous swellings below.

Vestiture: Mostly glabrous; a few scattered pale hairs dorsally.

Habitat: Fresh cow manure in pastures.

Remarks: The brown head, the three bare swellings at posterior end of grub,
and the epipharyngeal characters distinguish this (the largest) aphodian
larva.

TROX SUBEROSUS (Fab.)

Length 18 mm. Form elongate, slightly tapering posteriorly.

Width of head 3.8 mm.

Surface of head: Shining, rather coarsely but vaguely reticulate. Epicranial
stem a sharp narrow groove with raised edges. Epicranial arms impressed.
Anterior half of front with two vague longitudinal impressions near middle.

Color of head: Very deep chestnut brown.

Hpipharynx (fig. 3): Two-lobed. Arrangement of structures almost bilaterally
symmetrical. Details sufficiently shown in drawing.

Raster (fig. 6): An irregular transverse band, medially interrupted, of sparse,
blackish hairs.

Anal slit: Y-shaped; on each side a glabrous swelling with a fine black line
on it. ;

Vestiture: Each dorsal convexity of first six abdominal segments with a single
row of close, short, erect blackish spines. Entire grub with scattered dark
hairs, longest beneath.

Habitat: Soil under piles of old feathers, hair, or night soil.

Remarks: Together with the typical habitat, the peculiar epipharyngeal and
anal characters should make this grub easy to identify. - Each larva lives
in a vertical burrow down which he drags a wad of food material for
immediate needs.

b2 CIRCULAR 334, U. S. DEPARTMENT OF AGRICULTURE

PLECTRIS ALIENA (Chapin)

Length 28 mm. Form medium.

Width of head 3.5 mm.

Surface of head: Shining, rather coarsely, shallowly reticulate. Epicranial
stem a narrow impressed line through a Slight swelling. Postclypeal
margin of front with numerous fine punctures; above these two paramedian
pits; below curve of epicranial arm one large setose puncture and several
minute ones. Epicranial arms finely impressed.

Color of head: Dull yellow.

Epipharynx (fig. 8): Outline of labrum very angular, margins irregular.
Marginal spines coarse. Marginal striae distinct, long, sinuous and some-
times interrupted. Sensory eminence with numerous large and small
sensillae arranged in a transverse arcuate line. Five large fixed teeth.
Sensory hairs near tormae numerous and coarse. Tormae short and stout.

Raster (fig. 8) : Straight, pointed spines arranged in a large, spreading V made
up of 1 to 3 ranks of the inward-pointing spines. Outside of this about 3
irregular rows of larger, outward-pointing hooked spines. End and sides
of terminal segment with many rather long brown hairs, the former with
interspersed short stout spines.

Anal slit: Transverse, V-shaped.

Vestiture: A few scattered long hairs over entire surface of grub. The usual
small brown dorsal setae rather dense and conspicuous.

Habitat: Turf of golf courses, etc.

Remarks: A species of probably South American origin now found in the
South Atlantic States. While generally resembling common white grubs,
this larva is very different in details of structure, as shown in the drawings.

CHARACTERS DISTINGUISHING LARVAE OF POPILLIA, ETC.

Posterior end

Head-front view.
} of grub Width of head
| Anal slit 7] Epicranial
) Raster capuet
Third Epicranial
Legs; Second arm
< | First {emeest
Prothoraciny ASN VErER§ RE Siti
sclerotized JEeoS
plate Labrum
Huis : Mandible
oracic Spiracles Lower surface
segments E >¥ lof clypeus and
Ss ‘ w= \labrum to
a B show location
. of Epipharynx
Distal Sensory eminence
sclerotized plate
P Sensillae
I VABTARN Fixed spines
Articulated spines ie | iii Xs bi ie ake
| ; © . Zt
Lateral AL
ine ZB :
le Marginal
striae
Right
chs iNeak Left torma
Ciypeal .
Sclerotized x
plate Epigusta

Ciypeal sensillae

Popillia japonica Newm:

FieurR® 1.—A, diagram of a scarabaeid larva; B, diagram of head of a scarabaeid larva ;
C, epipharynx of Popillia japonica.

14 CIRCULAR 334, U. S. DEPARTMENT OF AGRICULTURE

Strigoderma arboricola (Fab) Pachystethus lucicola (Fab)

Pa

Macrodactylus subspinosus (Fab) Dichelonyx sp. Diplotaxis sp.

FIGURE 2.—Epipharynges of scarabaeid larvae.

| CHARACTERS DISTINGUISHING LARVAE OF POPILLIA, ETC. 15

“44}\
my

My

\

FE *®

=
=
Ss

fe Aphodius fossor(L) AA

Euphoria inda (L) Balint of
Ww

punctata(L) ,

FIGURE 3.—Epipharynges of scarabaeid larvae.

16 CIRCULAR 334, U. S. DEPARTMENT OF AGRICULTURE

Serica similis Lewis Serica paralleta Csy.

—— es tetleeemeee

= —_

Ochrosidia villosa(Burm)

Phyllophaga tristis (Fab.) Phyllophaga ephilida (Say)

FiGguRE 4,—Epipharynges of scarabaeid larvae.

CHARACTERS DISTINGUISHING LARVAE OF POPILLIA,,ETC. 17

Polyphylla 2 ANG kee .
variolosa (Hentz) a a 5

Xyloryctes
Sgiyrus (Fab)

FIGURE 5.—Epipharynges of scarabaeid larvae.

18 CIRCULAR 334, U. S. DEPARTMENT OF AGRICULTURE

\\\ aera Ahi
Ax Rea ? i oR
SSX LG Qs %
Se SE :
Zab) RS ce ck :
Ss =_ eae = :
Popillia Anomala AnetalG Strigoderma
Japonica Newm, — orientalis Waterh,

binotata Gyll. arboricola Fab

~
Ny

477
“il?
s7ttt >

4
NE
~
\
‘

¢ et eee
AZ TAN OF
"Wty,
‘ (EZ
WY
NANA.
WN

Pachystethus Macrodactylus

\
lucicola (Fab) subspinosus(Fab) | |

Phyllophaga

Phyllophaga
tristis (Fab.)

ephilida (Say)

rs . : first B
Diplotaxis sp. nd second legs second legs SY) YNend! of
+ ~ End of third le
Autoserica™” Mb: iihird leg second legs

castanea Arr. Serica similis Lewis End of third leg

Serica parallela Csy

Sly + \

SN Wy AZ
Sa UW GUY iG
\ NN Nee 1 has

SX j wT] TN Us
So} ey

er
Dichelonyx sp. Trox suberosus Fab.

Aphodius fossor(L) \
Figure 6.—Rasters of scarabaeid larvae.

CHARACTERS DISTINGUISHING LARVAE OF POPILLIA, ETc. 19

Ochrosidia Aphonus

villosa (Burm) castaneus (Melsh. Euphoria

3)
Qo.
re)
-
—

Osmoderma 1 ae on:
Carine nitida (L) eremicola Knoch Polyphylia variolosa (Hentz)

idnota - Xyloryctes
re encrate (L.) satyrus (Fab)

FIGURE 7.—Rasters of scarabaeid larvae.

20 CIRCULAR 334, U. S. DEPARTMENT OF AGRICULTURE

—_——_
/
-

raster .

Plectris.
aliena Chapin

FIGURE 8.—Epinharynx and raster of Plectris aliena.

U.S. GOVERNMENT PRINTING OFFICE: 1934

For sale by the Superintendent of Documents, Washington, D. C. - - - - - - Price 5 cents

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